Hunting and processing of straight-tusked elephants 125.000 years ago: Implications for Neanderthal behavior

Straight-tusked elephants (Palaeoloxodon antiquus) were the largest terrestrial mammals of the Pleistocene, present in Eurasian landscapes between 800,000 and 100,000 years ago. The occasional co-occurrence of their skeletal remains with stone tools has generated rich speculation about the nature of interactions between these elephants and Pleistocene humans: Did hominins scavenge on elephants that died a natural death or maybe even hunt some individuals? Our archaeozoological study of the largest P. antiquus assemblage known, excavated from 125,000-year-old lake deposits in Germany, shows that hunting of elephants weighing up to 13 metric tons was part of the cultural repertoire of Last Interglacial Neanderthals there, over >2000 years, many dozens of generations. The intensity and nutritional yields of these well-documented butchering activities, combined with previously reported data from this Neumark-Nord site complex, suggest that Neanderthals were less mobile and operated within social units substantially larger than commonly envisaged.

The well-preserved skull of this bull as well as its mandible have been very heavily restored for exhibition purposes, to such a degree that much of the original bone surface on these pieces is no longer visible. Apart from the skull and the attached cervical spine, Os sacrum with articulating vertebrae and Pelvis, most of the torso is missing. Table S6 compiles the qualitative and quantitative composition of bone complex E9 with 74 NISP representing 62 MNE. The following skeletal elements were studied for bone surface modifications: skull, mandible, complete cervical spine up to the 5 th cervical vertebra, thoracic vertebrae, lumbar vertebrae, sacrum, caudal vertebra 1, caudal vertebra 2, Scapula sin./dex., Humerus sin./dex., Radius/Ulna sin./dex., Femur sin./dex., Patella sin., Tibia dex., Fibula dex., Os carpi radiale dex., Os carpi intermedium sin./ dex., Os carpi ulnare sin., Os carpi acessorium sin., Os carpale I sin./dex., Os carpale II dex., Os carpale III sin./dex., Os carpale IV dex., Os metacarpale I-IV sin./dex., Talus dex., Calcaneus sin., Os tarsale III dex., 3 Phalanges proximales sin., Phalanges indet. Of these, an MNE of 8 shows anthropogenic traces on elements of the extremities. Table S7 lists the observed cut marks as a result of different activities during the butchering process. These testify to the systematic separation of the connections between the zonoskeleton, stylo-, zeugo-, and autopodium. Traces appear repetitively as can be seen, for example, on the right and left Femora. Remarkable is also abundant evidence for disarticulation of hand and foot bones.
In addition to cut-marked bones, three skeletal elements show traces of gnawing by a medium-sized carnivore. These were observed on the proximal right Scapula, on a lumbar vertebra, and the Os metacarpale II and III sin., and produced when after butchering these bones were still in articulation. Carnivore modification did not lead to the disarticulation of the metacarpals ( Figure S8). Bone complex E10 is described by Mania as consisting of the primary context remains of a large individual that had been disarticulated by humans and was in an advanced stage of decay. The individual body parts were distributed over a larger area, of 5x6 meters, and the anatomically correct position of particular body parts was still recognizable. Mania notes the absence of the skull and some long bones, elements that were not discovered even after subsequent spatial expansion of the excavation area. The complex is characterized by an almost complete torso in anatomically correct position ( Figures S9-S10). The anterior part of the carcass with the cervical spine, including the first two cervical vertebrae, parts of the torso, and the anterior limbs lay eastwards, at a distance of 3 meters. The complete left leg in an anatomically correct position lay on the cervical spine (23). It is worth remarking that below the thorax of this animal, some of the gut content was still preserved (59). The animal suffered from hyper-ossification of the sacrum due to arthritis (3).
Later paleontological studies showed that bone complex E10 comprises the remains of altogether 3 individuals: E10A, a 47-year-old male whose average body mass was calculated to be 12,5 tonnes with a shoulder height of 395 cm, E10B, a 26-year-old female with a shoulder height of more than 3 meters and an estimated body weight of more than 5,5 tonnes and an indeterminate additional individual (E10C) (3,24). E10A is represented by a vertebrate column with most of the ribs and the Pelvis, nearly complete left and right forelegs with only the right Scapula, and nearly complete left hindleg with only the left Femur preserved. Larramendi et al. (1) reconstructed the female E10B based on the Os sacrum and a nearly complete additional Pelvis.
Individual E10B was identified through the presence of a complete Pelvis with Os sacrum. Individual E10C was defined due to the presence of a 5th cervical vertebra, Tibia dex., Os carpi intermedium indet., Os carpale IV dex. and an Os metatarsale III dex..
The carcass of individual 10A survived in large parts, in contrast to individual 10B, only represented by a Pelvis with attached Os sacrum. Due to the additional presence of a Tibia and some autopodium bones and a cervical vertebra, the third individual within the bone complex was identified.
In total 48 specimens (38 MNE) showed signs of anthropogenic modification. Table S9 lists the observed cut marks as a result of different activities during the butchering process. The detailed analysis of the cut marks shows that the carcass was dissected into its smallest parts. The butchery sequence shows acquisition of the tenderloin and the dissection of the thorax. Cut marks on the midshafts of the right 14th-16th and 18th ribs could indicate that the inside of the intact ribcage was treated with stone implements after the torso was broken open, the viscera removed and the carcass rested on the right side. On the ribs of the left half of the body, the cut marks are lateral, e.g., on the midshafts of the 6th, 7 th , and 16th ribs. If one follows the above interpretation, one would have to postulate a position of the torso on the left side. Cut marks on the lateral surfaces of the spinae of vertebrae, which were created during the removal of the tenderloin, were probably also created while the torso rested on the left side.
The deboning, defleshing and systematic dissection of the skeleton into its individual components of the right and left halves and the evisceration of the carcass are attested. It is remarkable that here, as in the previous examples, the dismemberment of the hands and feet is evident and that none of the skeletal elements was opened for the removal of bone marrow.
The Pelvis with attached Os sacrum attributed to individual 10B also shows cut marks. They indicate the deboning of the right hind leg for this individual. Table S10 lists the observed cut marks as a result of different activities during the butchering process. For E10C none of the bones were cut marked.
The bones of individual 10A show not only anthropogenic traces but also those of modification by medium-and large-sized carnivores. A total of seven bones are affected. The spinae of two thoracic vertebrae, one of them with cuts, show slight traces of gnawing proximally, as does the distal end of a rib. In addition, the Os metacarpus II sin., the Phalange proximale, dig. III man. sin. and the Os metacarpus V dex. are punctured distally. Finally, on the right Tibia, which was already mentioned in context with the cut marks, traces of gnawing can be seen at the proximal end. Distally, the bone shows a modern fracture.

Unit 6.1 -lower littoral zone (PAZ IV of the Eemian Interglacial)
Bone complex E23 Bone complex E23 was excavated from the southern part of the main find horizon, the lower littoral zone (unit 6.1), located next to complexes E21 and E24. Mania assumes that all three originally formed one large complex (23). Remains from altogether three individuals were scattered over an area of 6x6 meters. E23A, a large male, 44 years of age with an estimated body mass of 9,8 tonnes and a shoulder height of 363 cm, was associated with remains of an additional, even larger, 50 years old male individual of 13 tonnes and a shoulder height of 400 cm (E23B), as well as a small female, younger than 26 years, with an estimated 2030 kg body mass (E23C) (1,24). This bull, E23B, represents one of the largest fully grown individuals excavated at Neumark-Nord 1.
Concerning the find-situation ( Figures S13-14), the anatomical order of body parts was still recognizable though not in anatomical connection. The bull (E23A) lacked its skull, but Atlas and Os hyoideum with ossified traumatic fracture (3,24) as well as some cervical vertebrae were distributed all over the area (23). The Pelvis, spine and ribs were more or less in anatomical position. Only a few vertebrae were in direct anatomical association. Mania counted 26 ribs and 27 vertebrae, excluding the tail (23). The Scapula was discovered at a distance of 4 meters. Skulls, tusks and long bones were also scattered over the area covered by the bone complex (23). Bones studied for bone modifications comprised a NISP of 486 (95 MNE) ( Table S11). The following elements were studied for Individuals E10A-10C: Cranium with M3 sup. dex. and tusk, Mandible, Os hyoideum dex., Atlas and cervical vertebrae 3-7, thoracic vertebrae 1-6, 8-9, 11-16, 18-19, lumbar vertebrae 3-4, caudal vertebrae 1, 2 and 4. From the right side of the rump rib 1, 3-6, 8-14, 16-17 survived, from the left side rib 1-2, 5-9, 12-17, 26 additional rib fragments and the Scapula survived. The foreleg is represented by Humerus dex. and Radius sin., Os carpi intermedium sin., Os carpale II and III sin., Os carpale III dex., Phalanges proximales, dig. III-IV, Phalanges distales, dig. III-IV. The right hindquarter with Os sacrum, Pelvis, Femur, Fibula and Calcaneus was recorded in addition to Os metatarsale I sin.. Cut marks were recorded on 34 NISP, representing an MNE of 33 (Table S11). The majority of cut marks were observed on vertebrae and ribs of individual E23A. These traces testify to the complete dismemberment of the torso. The exploitation of the tenderloin is evident here as well as the removal of skin, fat and the connecting tissue between the ribs. This process was meticulously carried out from the outside of the carcass but also from within the area of the inner rib cage. In addition, marks refer to the disarticulation of the entire spine and rib cage. Traces on long bones prove the separation of the stylo-, from the zeugo-, from the autopodium after the right front leg and the left hind leg have been deboned. Finally, cuts on bones of the left metacarpus indicate its disarticulation.
Carnivore traces were additionally observed on bones from bone complex E23. From individual E23A, the cervical spine is more or less completely preserved ( Figure S16). The bones show gnawing marks on the right side of the transverse process. The character of the marks suggests that the cervical spine was still articulated at the time when carnivore gnawing occurred. The same is true for the distal ends of a 3rd, 5 th , and 6th rib, which show carnivore tooth marks.

Unit 7 (PAZ V, the Carpinus phase of the Eemian Interglacial)
Bone complex E30 Bone complex E30 was located in the eastern part of the lake basin, in the lower part of the upper "gewarvte Feindetritusmudde" sensu Mania (23). He (23) describes bone complex E30 as containing the remains of a large adult elephant, which probably became embedded more or less completely. Although the skull was preserved in numerous fragments, the tusks were missing. The fragment of a flint flake was uncovered among the bones ( Figure S17). On its dorsal and ventral sides, remains of crushed oak bark were identified, originally interpreted as remains of a hafting residue, an interpretation disputed by Koller and Baumer (60) who suggest that the crushed oak bark may have been used for the tanning of hides. Cut marks were observed on 6 bones, constituting 5 skeletal elements. Even though not many anthropogenic traces were observed, the traces present indicate the disarticulation of major parts of the carcass. Particularly remarkable are cut marks observed on the condyles of the occipital, next to the opening of the occiput. These marks are particularly deep, cross each other and are confined to a small area. They suggest that the severing of the skull was done by the disarticulation at the atlas. Cut marks on the Scapula show that the right half of the carcass was dissected and traces on ribs indicate the disarticulation of the ribcage and the removal of fat and connective tissue between the ribs (Table S14). Finally, as observed in all complexes illustrated here, the feet were disarticulated as a means to harvest the rich fat resources stored in the elephant's foot cushion. Traces of carnivore modification were not observed on bones from E30.

Supplementary Text 2: Calculating estimates of caloric and nutritional yields of P. antiquus from Neumark-Nord 1
Given the very large sizes of the Neumark-Nord straight-tusked elephants and their extensive processing, they undoubtedly yielded large amounts of food. Based on studies of contemporary elephants it is possible to set some constraints on the nutritional yields provided by these elephants.
African elephant carcasses average 38% muscle tissue/weight, with little variation, and including brain, tongue, liver, heart, and trunk, so the edible mass (omitting stomach content, etc.) makes up 42% of body weight (61), a more generous estimate than an earlier one by Frison and Todd (62) (25-35%). For a 10-tonne Neumark-Nord male elephant this implies 4200 kg of edible material.
However, there are constraints on the consumption of meat, which should be taken into account when discussing the food yielded by the Neumark-Nord elephants. In general, mammal tissue is rich in proteins, relatively low in fats (although high amounts are stored in bone marrow and adipose tissue), and contains almost no carbohydrates. An unbalanced intake of these macronutrients, in the form of a diet without sufficient non-protein calories (i.e. calories from either fat or carbohydrates), can lead to severe health problems including a condition known as "rabbit starvation" (37,38). Protein should generally not exceed 25-35% of the daily macronutrient energy intake (en%) (38,63), a proportion which is reflected in hunter-gatherer diets (37,38,64,65). A Neumark-Nord elephant carcassas well as those of all other mammals -could therefore come with a considerable excess of protein, when all edible parts are considered.
The body fat percentage of the Neumark-Nord elephants is unknown, but studies of current elephants shed light on the range of likely values. Studies of captive elephants have documented body fat percentages of 5.24 to 15.97% for Loxodonta africana (66), and a range of 3.54 -24.59% for Asian captive elephants, Elephas maximus (67).
Based on this we can suggest the likely energy yield in fat for the elephants in this study. For a 10-tonne Neumark-Nord male elephant, a conservative estimate of 3.5% body fat would yield minimally 350 kg of fat, probably the lower limit of variation throughout a year. Based on Byer's and Ugan's (61) calculations as well as the body fat percentages discussed above, a 10-tonne elephant would yield 4200 kg of edible material, including minimally ~350 kg of fat. With 100 g fat translated into ~660 kcal (33) that would imply that our Neumark-Nord male would have stored minimally 2.3 million kcal in the form of fat only. Continuing this thought exercise, and assuming a high daily energy expenditure (DEE) of 4000 kcal/day for an "average" Neanderthal ((42), p. 326-27) we are talking about minimally 575 daily rations financed only by the lipids coming from one large elephant.
Further, taking the 'protein ceiling' into account, we can estimate the amount of meat and fat that would be available for consumption. Following Speth's calculations (38), and assuming our average Neanderthal would have an upper protein intake level of 300 g/day at 4 kcal/g and meat consisting of 20% protein, this would imply a daily safe maximum intake of 1500 grams of "meat", yielding 1200 kcal. Combining the meat plus lipids into a 4,000 kcal "protein safe" elephant-fat and -meat package would yield >821 daily rations, a very conservative estimate, based on the lowest values of the documented body fat range. In such an "elephant-products only" scenario, i.e. without alternative sources of key macronutrients, about 3,000 kg of "meat" would not be usable. Increasing the body fat content to 10% (a value which is still in the middle of the range calculated for contemporary African elephants) would yield about 2,500 "elephant-only" daily portions in our calculation, with a good 1,000 kg of muscle meat still unused. Raising the fat content of our hypothetical elephant to 12.8%, still below the middle of the range of Asian elephants, would enable complete and safe consumption of this elephant, yielding >3,000 adult Neanderthal rations of 4,000 kcal. This is of course a thought exercise rather than an exact value, which is intended to illustrate the quantity of food yielded by the Neumark-Nord elephants, even taking nutritional constraints into account.
Edible material from elephants was likely supplemented with lipids obtained from other animals, e.g. by marrow-processing activities documented at the nearby location of Neumark-Nord 2 (17), or e.g. with carbohydrates from plant foods. While the processing of large herbivores created a clear archaeological signal in the Neumark-Nord landscape and at Neanderthal sites in general, nutritional studies indicate that they could not have survived on terrestrial game alone and that plants played an important role in Neanderthal diets, providing carbohydrates and some of the required nutrients and calories (68,69). Recent studies suggest that starch-rich foods were already important before the split between the Neanderthal and modern human lineages (70), and document consumption of a similarly wide range of plant species by Neanderthals across their geographical range (69,71). The plant component of the Neanderthal diet at Neumark-Nord remains largely unknown, as is the case at most Palaeolithic sites, no matter how important plant foods must have been in the past, especially in the context of the protein ceiling. At Neumark-Nord, fragments of charred hazelnut (Corylus avellanus), acorn (Quercus sp.), and blackthorn/sloe plum (Prunus spinosa) (72) may constitute the ephemeral traces of this part of Last Interglacial Neanderthal diet (22).

Fig. S1
Topographic map of the basin Neumark-Nord 1 (center) and Neumark-Nord 2 (upper right) with the spatial distribution of bone complexes (dots) mentioned in the SI from units 4, 6.1, 6 and 7 (after Mania (23) and archeological rescue interventions (black rectangles), the highdensity distributions of flint artifacts and fragmented faunal remains (hatched areas), and the position of the Neumark-Nord 2 excavated area (in black) (after (22). Contour lines of the basins, at respectively 8, 16, and 24 m below the surface. The outer line indicates the maximum extension of the water body.  Table S5.

Fig. S4B
Cut marks on Os carpi intermedium sin., proximal aspect. For a detailed description see Table  S5.  Table S7.

Fig. S7B
Bone complex E9. Cut marks on Femur dex.distal / caudal aspect. For a detailed description see Table S7.

Fig. S7C
Bone complex E9. Cut marks on Femur sin.distal / caudal aspect. For a detailed description see Table S7.

Fig. S7D
Bone complex E9. Cut marks on Talus dex.cranial aspect. For a detailed description see Table S7.

Fig. S7E
Bone complex E9. Cut marks on Os carpale IV dex.distal aspect. For a detailed description see Table S7.

Fig. S11B
Bone complex E10, Individual E10 A. Cut marks on 7 th rib -lateral aspect. For a detailed description see Table S9.

Fig. S11C
Bone complex E10, Individual E10 A. Cut marks on Pelvis dex. -Os ilium -medial aspect. For a detailed description see Table S9.

Fig. S11D
Bone complex E10, Individual E10 A. Cut marks on Scapula dex. -medial aspect. For a detailed description see Table S9.

Fig. S11E
Bone complex E10, Individual E10 A. Cut marks on Humerus dex.distal/cranial aspect. For a detailed description see Table S9.

Fig. S11F
Bone complex E10, Individual E10 A. Cut marks on Radius/Ulna sin. -distal aspect. For a detailed description see Table S9.

Fig. S11G
Bone complex E10, Individual E10 A. Cut mark on Os carpi intermedium dex. -proximal aspect. For a detailed description see Table S9.

Fig. S11H
Bone complex E10, Individual E10 A. Cut marks on Os carpi intermedium sin. -medial aspect. For a detailed description see Table S9.

Fig. S11I
Bone complex E10, Individual E10 A. Cut marks on Os carpi ulnare sin. -proximal aspect. For a detailed description see Table S9.

Fig. S11L
Bone complex E10, Individual E10 A. Cut marks on Os metacarpale III dex. -proximal aspect. For a detailed description see Table S9.

Fig. S11M
Bone complex E10, Individual E10 A. Cut marks on Os metacarpale IV sin.distal/dorsal aspect. For a detailed description see Table S9.

Fig. S11O
Bone complex E10, Individual E10 A. Cut mark on Tibia dex.prox. Diaphysiscranial aspect. For a detailed description see Table S9.

Fig. S11P
Bone complex E10, Individual E10 A. Cut marks on Talus dex.cranial aspect. For a detailed description see Table S9.  Table S9.  Table S12.

Fig. S15G
Bone complex E23. Cut marks on Humerus dex.proximal -dorsal aspect. For a detailed description see Table S12.

Fig. S15H
Bone complex E23. Cut marks on Radius / Ulna sin.distal aspect. For a detailed description see Table S12.

Fig. S15I
Bone complex E23. Cut marks on Os sacrum -lateral aspect. For a detailed description see Table S12.

Fig. S15K
Bone complex E23. Cut mark on Femur dex.distal -caudal aspect. For a detailed description see Table S12.   Bone complex E30. Cut marks on left (above) and right (below) Condylus occipitalisdistal aspect. For a detailed description see Table S14.

Fig. S18B
Bone complex E30. Cut marks on Os tarsi centrale dex.distal aspect. For a detailed description see Table S14.

Table S1
P. antiquus individuals from Neumark-Nord 1 studied for bone surface modifications. Data for composition of bone complexes for Unit 6 and 6.1 and sex, age and body mass taken from (1,23,24). C1-C3 were not provided with designations for bone complexes. MNI -Minimum number of individuals, NISP -Number of identified specimens per taxon, MNE -Minimum Number of Elements, NISPcut / MNEcut -Number of identified specimen per taxon with cut marks / Minimum number of cut marked elements, NISPgnaw / MNEgnaw -Number of identified specimens per taxon modified by large carnivores / Minimum number of elements modified by large carnivores.

Table S2
The studied sample, listed according to skeletal element. NISP -Number of identified specimens per taxon, MNE -Minimum Number of Elements, NISPcut / MNEcut -Number of identified specimens per taxon with cut marks / Minimum number of cut marked elements, NISPgnaw / MNEgnaw -Number of identified specimens per taxon modified by large carnivores / Minimum number of elements modified by large carnivores.

Table S4
Number of identified specimens per taxon (NISP), Number of Elements (MNE) and Number of identified specimens per taxon with cut marks (NISP cut) and Number of Elements with cut marks (MNE cut) for bone complex E6.

Table S5
Cut marks and sequences of disarticulation observed for bone complex E6.

Table S6
Number of identified specimens per taxon (NISP), Number of Elements (MNE) and Number of identified specimens per taxon with cut marks (NISP cut) and Number of Elements with cut marks (MNE cut) for bone complex E9.

Table S7
Cut marks and sequences of disarticulation observed for bone complex E9.

Table S8
Number of identified specimens per taxon (NISP), Number of Elements (MNE) and Number of identified specimens per taxon with cut marks (NISP cut) and Number of Elements with cut marks (MNE cut) for bone complex E10.

Table S9
Cut marks and sequences of disarticulation observed for bone complex E10, Individual E10A.

Table S10
Cut marks and sequences of disarticulation observed for bone complex E10, Individual E10B.

Table S11
Number of identified specimens per taxon (NISP), Number of Elements (MNE) and Number of identified specimens per taxon with cut marks (NISP cut) and Number of Elements with cut marks (MNE cut) for bone complex E23.

Table S12
Cut marks and sequences of disarticulation observed for bone complex E23.

Table S13
Number of identified specimens per taxon (NISP), Number of Elements (MNE) and Number of identified specimens per taxon with cut marks (NISP cut) and Number of Elements with cut marks (MNE cut) for bone complex E30.

Table S15
The Neumark-Nord 1 elephant bone complexes in their geological and archaeological setting of the Neumark-Nord 1 and 2 sequences, with indicated Pollen Assemblage Zones (PAZ) of the Last Interglacial (25,58) for Neumark-Nord 1 and 2, the various lake phases of Neumark-Nord 1 (tr = transgression, re = regression), the sedimentary context of the various find levels as well as information on the associated faunal and lithic finds (compiled after (9, 16-20, 22, 29, 73) The provenance of elephant bone complexes/assemblages analyzed is presented in a separate column (see also Table S1).